FINAL REPORT: Macroinvertebrate Inventory of the Caspar Creek Watershed

Kenneth W. Cummins1 and David Malkauskas2

1. California Cooperative Fisheries Research Unit, Humboldt State University 2. Department of Entomology, Michigan State University

1 Introduction

A macroinvertebrate assessment was conducted on the Caspar Creek Experimental Watershed in northwestern California using a functional feeding group (FFG) classification. The approach, developed over 30 years ago (Cummins 1973), has been tested, modified, and employed in many studies (e.g. Cummins and Klug 1979, Cummins and Wilzbach 1985), Merritt and Cummins 2006). The approach categorizes macroinvertebrates based on their morphological and behavioral mechanisms by which they acquire one or more of six general food types: coarse particulate organic matter (CPOM), fine particulate organic matter (FPOM), periphytic, non-filamentous algae and associated bio-film, invertebrate prey organisms, and filamentous algae (Table 1). The morpho-behavioral adaptations of stream macroinvertebrates for acquiring food are easily observed in the field on live specimens. An example is the large eyes, bright color patterns, and active movement that characterize the three predaceous stonefly families (setipalpian Plecoptera). The abundance (especially expressed as biomass) of any FFG is an indicator of the relative availability of its food resource category. The method has the advantage that the survey crew leaves the field with the data as well as with preserved samples that can be analyzed in taxonomic data, and measured for biomass conversion calculations in the lab, if (and this is almost always the problem) funds are available for the very time-consuming process of microscope analysis. The point is, that a great deal of information about the status of a stream ecosystem can be obtained in the field using the FFG approach, but it does not for close the ability to analyze the samples in the lab.

The field numbers can be converted to biomass using length-mass regressions, or to calculate any of the commonly used indices of stream condition (e.g. % Ephemeroptera, Plecoptera, and Trichoptera in a sample). Ecological tables in Merritt et al. (2008) provide information of the functional feeding group categorization of essentially every genus of North American aquatic insect. Information in Pennak (1989) can be used to assign non-insect invertebrates to functional groups.

Methods

The FFG field approach was used to rapidly characterize the North and South Branch tributaries and the Mainstem of Caspar Creek with regard to a number of ecosystem attributes of each. Field collections, together with laboratory analysis, were used to compile a taxonomic list of stream invertebrates for Caspar Creek. The samples taken in May and October of 2008 were compared to those taken by Bottorff and Knight (1996) from 1986 through 1994 to determine if taxonomic changes had occurred (Appendix 1). Timed 30 second samples were taken with a D-frame dip net (250µm mesh) in three habitats: riffle (coarse sediments), pools and backwaters (fine sediments), and plant litter accumulations (drop and trap zones). Three samples were taken pseudo- randomly (i.e. haphazardly) in each habitat type at locations along a 50 meter reach of stream at five locations. The sites were: upper and lower North Fork, lower South Fork, and upper and lower Mainstem (Fig. 1). The samples were sorted fresh at stream side primarily with the unaided eye but with some small specimens checked using a 5X hand lens. The

2 invertebrates were sorted into functional groups (Table 1) and the numbers in each discernable taxon recorded. A small portion of a sample was examined at a time in a white enamel tray. After sorting a tallying the invertebrates, the sample was placed in a Whirlpac™ bag, labeled, and preserve with 75% ethanol for return to the lab. Large mineral substrates and pieces of wood were thoroughly washed and discarded. The fine sediments were placed in a bucket agitated and repeatedly decanted onto a 250µm mesh sieve until no more light materials were visible in the wash. The litter samples were examined piece by piece, small portions at a time. The average processing time for a field sample was 1 to 2 hours. In the laboratory, each sample was examined under a dissecting microscope and specimens were identified (using Merritt et al. 2008 and Pennak 1989) and measured to the nearest mm. The number of specimens of a given taxon in each mm size category was used to convert the numbers to dry biomass using the INVERTCALC program (Cummins, Merritt and Kolouch, Humboldt State University). Each taxon is coded, allowing the data on that taxon to be tabulated by functional group.

Once the data had been summarized, a number of ratios of functional groups are used as surrogates for selected stream ecosystem attributes (Table 2). The ratios were calculated for the three habitats for each of the 5 sites for the fall samples (Table 3). Because the major analysis utilized dimensionless ratios, the resulting values are fairly independent of sample size. Particular emphasis was placed on the characteristics of the riparian vegetation cover at each sample site, i.e. percent shading and relative proportion of evergreens to deciduous trees.

Results

Using the proposed threshold values for the functional group surrogate ratio for the autotrophic/heterotrophic (P/R) index (i.e. > 0.75 = autotrophic, Merritt et al. 1996, 2002, 2008). This ecosystem attribute(P?R) is directly measured as the ratio of gross primary production to total community respiration. The Caspar Creek water shed is heterotrophic (Table 3). This is as expected because the stream is heavily shaded throughout the drainage and nearly canopy closed along the North and South Forks and the Upper Mainstem. The riparian is dominated by second growth conifers and the colonizing hardwood red alder. The South Fork is bordered by relatively more red alder, but the entire drainage has a mix of evergreens and deciduous hardwoods. The expected shredder index (Table 2), which reflects the abundance of coarse organic litter (CPOM) relative to fine organic litter (FPOM), was well above 0.50 at all sites in October. As expected, the values were highest in the litter samples, but since coarse litter can be trapped or deposited in any of the habitats, shedders were wildly distributed. Therefore, taken together, the macroinvertebrate surrogate ratios for P/R and CPOM/FPOM (Table 3) characterize Caspar Creek as an heterotrophic ecosystem with a litter supply that supports a strong linkage between the riparian vegetation and shredder macroinvertebrates. If the presence – absence taxonomic data from Botteroff and Knight (B and K, 1996) are compared to that for the present study (PS) in 2008 (Appendix Table) for the functional group surrogate P/R ratio, both indicate Caspar Creek is heterotrophic (i.e. < 0.75). The ratios are essentially the same: B and K = 0.33 and PS = 0.35. Similarly, the surrogate for

3 CPOM/FPOM indicates a normal linkage between shredders and riparian litter for an heterotrophic system (i.e. > 0.50; B and K = 0.59, PS = 0.69). This is in agreement with the sample data from the present study (Table 3) in which the surrogate ratios ranged from 0.78 to 30.40 for the five sites. The ratio of suspended FPOM to benthic FPOM as indicated by the functional group surrogate (Table 3) is below or at the threshold value of 0.25 (B and K = 0.15, PS = 0.26). This lack of suspended FPOM in the system is likely a partial explanation for the poor representation by filtering collectors in Caspar Creek (Table 3, Appendix Table). Not only the quantity of FPOM in transport but also its quality as a food resource could be limiting the filtering collectors. The availability of stable substrates, i.e. coarse sediments in riffles, large wood, and rooted aquatic vascular plants, as measured by the functional group surrogate, indicates they are scarce (threshold = 0.5, Table 2; Appendix Table, B and K = 0.45, PS = 0.53). This is reflected in the relative scarcity of large wood, poorly represented riffles, and the scarcity of rooted plants in Caspar Creek. The scarcity of stable substrates is likely also a factor in the low abundance of filtering collectors in Caspar Creek (Table 3, appendix Table), because they require such locations to set up their filtering stations. The ratio of behavioral drifting macroinvertebrates to accidental drifters is an indicator of the reliable dawn and dusk food supply for macroinvertebrates in the water column to serve as prey of drift-feeding salmonids. In Caspar Creek in October, less than half of the macroinvertebrate fauna was represented by behavior drifting taxa ( Table 3, 21 to 37%) indicating possible less than optimal food supply for drift-feeding salmonids.

Taxonomic differences in the macroinvertebrate fauna reported by Bottorff and Knight (1996) and the present study were not great indicating that Caspar Creek has not changed significantly in the ensuing 12 years. The differences observed (Appendix 1 can be attributed to a number of causes. 1) The study by Bottorff and Knight (1996) was conducted over 8 years and the longer time frame and greater number of samples increased the probability of collecting rarer taxa, for example the large megalopteran predator Dysmicohermes. 2) Changes in the taxonomy resulting over the 12 years, for example Bottorff and Knight (1996) used Merritt et al. (1996, 3rd edition) and the present study used Merritt et al. (2008, 4th edition). This is reflected in changes in groups such as the mayflies and caddisflies. 3) The use of artificial substrates by Bottorff and Knight (1996), namely rock baskets and leaf packs, increased the probability of attracting and concentrating riffle forms and litter-feeding taxa (shredders). 4) Sampling with a D-frame net used in the present study allowed for the collection of surface skating forms, such as Gerridae and Gyrinidae, which is unlikely when artificial substrates are used because the surface skaters are lost when the substrates are removed from the water. 5) A major part of the difference is the greater taxonomic resolution employed I the present study in the dipteran family Chironomidae. In Bottorff and Knight (1996) all representatives of the family were lumped together, while in the present study the specimens were sorted to subfamily and genera. (Appendix 2). This also accounts for some of the differences in functional group totals (Appendix Table) between the two

4 studies because the Chironomidae is a large group representing several functional groups. Whereas Bottorff and Knight (1996) categorized all specimens as collector-filterers.

In summary, Caspar Creek can be characterized as being in a stable period with regard to the macroinvertebrate fauna, and by extension using the functional group surrogate ratios, also in a stable period considering selective ecosystem attributes (Table 3, Appendix 1). Caspar Creek has been and still is heterotrophic with a stable linkage between shredders and the riparian vegetation that borders the stream and provides their litter food supply. Of the four taxa that occurred at all five sites, two are shedder-detritivores (Hydatophylax and Lepidostoma) and two are collector-gatherers (Paraleptophlebia and Oligochaeta). Stable substrates are marginally available and the FPOM food supply for collector- filterers is poor. This has consequences for the drift-feeding salmonids because many of the filtering collectors are behavioral drifters that would be reliably available to dawn and dusk drift-feeding fish. In general, Caspar Creek invertebrate taxa do not represent a dominance of behavioral drifters (Table 3). Future harvest plans, if they include alteration of the riparian zone could have significant impact on the macroinvertebrate communities and, by extension, the resident salmonids of Caspar Creek.

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5 Table 1. Macroinvertebrate functional feeding group (FFG) characteristics. CPOM = coarse particulate organic matter, FPOM = fine particulate organic matter. (Data combined and modified from Merritt et al. (2008).

Functional feeding Primary food Secondary food Voltinism (no. group resource resource for generations / y) facultative taxa Shredders

Plant litter CPOM Microbially Some facultative for Shredders conditioned riparian FPOM Univoltine, a few plant litter semi-voltine

Herbivore Live vascular None (obligate for shredders aquatic plants primary food resource)

Scrapers Non-filamentous Some facultative for Univoltine (some periphyton algae FPOM bivoltine) and associated biofilm

Filtering collectors FPOM in A few facultative Polyvoltine or suspension scrapers bivoltine

Gathering collectors Deposited FPOM A few facultative Polyvoltine or scrapers bivoltine

Predators Engulfing forms Prey (tend to be smaller) Some facultative for Usually semivoltine FPOM in first and (some univoltine) Piercing forms Prey (tend to be second instar larger)

Piercers (herbivores Single cells of None (all obligate Univoltine only present with filamentous algae for primary food filamentous algae) resource)

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Table 2. Examples of functional feeding group ratios serving as surrogates for stream ecosystem attributes. Modified from Merritt et al. (2008).

Attribute Direct measurement of Functional feeding Suggested thresholds for ecosystem attribute group surrogate FFG attribute ratios from previous studies

Ratio of autotrophy to Gross primary Scrapers + heribovre Autotrophic system ≥ heterotrophy (P/R) production as a ratio of shedders as a ratio of 0.75 total community plant litter shredders + respiration (i.e. oxygen total collectors consumed, including by primary producers

Riparian linkage to Leaf pack studies with Plant litter shredders as Expected linkage shredders riparian species utilizing a ratio of total collectors between riparian data on percent cover of (CPOM vegetation and shredders those riparian species consumers/FPOM Fall-winter shredder consumers) populations > 0.50 Spring-summer shredder populations > 0.25

Habitat stability for Inventory of areal Filtering collectors + Adequate stable macroinvertebrates coverage of stream scrapers (+herbivorous substrates > 0.50 bottom by boulders, shredders if present) as cobbles, bedrock, and a ratio of total shredders LWD + gathering collectors

Annual flow regime Annual and longer flow Semivoltine invertebrate Predictable annual flow records (recurrence taxa as a ratio of regime > 0.10 intervals) polyvoltine invertebrate taxa

Top-down predator Gut analyses of Predators as a ratio of Expected predator-prey control of invertebrate predators and total of all other balance = 0.10 – 0.20 community structure determination of density functional feeding and generation times of groups prey in guts

Quantity and quality of Quantity and quality Filtering collectors as a Expected quantity and FPOM in transport (O2 consumption or loss ratio of gathering quality (i.e. sufficient to (organic portion of of mass on ignition) of collectors support suspension suspended load that can FPOM in transport as a feeders) of FPOM in serve as a food resource ratio of CPOM and transport > 0.25 for suspension feeders FPOM in storage in and on the sediments

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References

Cummins, K.W. 1973. Trophic relations of aquatic insects. Annual Review of Entomology 18: 232-276.

Cummins, K.W. and M.J. Klug. 1979. Feeding ecology of stream invertebrates. Annual Review of Ecology and Systematics 10: 147-172.

Cummins, K.W. and M.A. Wilzbach. 1985. Field procedures for the analysis of functional feeding groups in stream ecosystems. Appalachian Environmental Laboratory, Contribution No. 1611, University of Maryland, Frostburg, MD.

Merritt, R.W., J.R. Wallace, M.J. Higgins, M.K. Alexander, M.B. Berg, W.T. Morgan, K.W. Cummins, and B. VandenEeden. 1996. Procedures for the functional analysis of invertebrate communities of the Kissimmee River-floodplain ecosystem. Florida Scientist 59:216-274.

Merritt, R. W., K. W. Cummins, M. B. Berg, J. A. Novak, M. J. Higgins, K. J. Wessell, and J. L. Lessard. 2002. Development and application of a macroinvertebrate functional groups approach in the bioassessment of remnant oxbows in the Caloosahatchee River, Southwest Florida. JNABS 21:290-310.

Merritt, R.W. and K.W. Cummins. 2006. Trophic relationships of macroinvertebrates. Pages 585-609 in Hauer, F.R. and G.A. Lamberti (eds.), Methods in stream ecology, 2nd edition. Academic Press, San Diego.

Merritt, R.W., K.W. Cummins, and M.B. Berg (eds.). 2007. An introduction to the aquatic insects of North America, 4th edition. Kendall/Hunt Publishers, Dubuque, IA. 998 p.

Pennak, R. W. 1989. Freshwater invertebrates of the United States. 3rd edition. John Wiley and Sons, Inc., N.Y. 628p.

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Appendix 1. Comparison of the taxa collected by Bottorff and Knight (1996; North Fork) and in the present study (2008; North and South Forks and Mainstem) in Caspar Creek, Jackson State Forest, CA. (X = present in samples; * = Collected in only one of the studies; Shred-detrit = shredder-detritivore; Shred-herb = shredder-herbivore; Coll-gather = collector-gatherer; Coll-filt = collector-filterer; Scrape = scraper; Pred = predator; Piercer = macrophyte piercer. Taxa Bottorff Functional Present Functional and Groups Study Groups (Merritt Knight (Merritt and and Cummins Cummins 2008) 1984) Collembola (springtails) X Coll-gather X Coll-gather Entomobryidae X Coll-gather Hypogasturidae X Coll-gather Simithuridae X Coll-gather; (facultative Shred- herbivore) Tomoceridae X Ephemeroptera (mayflies) Siphloneuridae now Ameletidae Ameletus X Coll-gather X Coll-gather; A. cooki (facultative Scrape) Baetidae Baetis X Coll-gather X Coll-gather B. bicaudatus B. tricaudatus Diphetor hageni X Coll-gather ? Fallceon quilleri X Coll-gather ? Ephemerellidae Drunella X Scrape X Scrape (facultative pred) D. flavilinea X Scrape Ephemerella X Coll-gather X Coll-gather; E. dorothea infrequens (facultative Scrape) Serratella X Coll-gather X Coll-gather

S. teresa S. tibialis Timpanoga hecuba X Coll-gather X Coll-gather

Heptageniidae Cinygma X Scrape X Scrape;

9 (facultative Coll- gather)

Cinygmula X Scrape X Scrape; (facultative Coll-gather) Epeorus X Coll-gather X Scrape; (facultative Coll-gather)

Ironodes X Scrape X Scrape; (facultative Coll-gather) Nixe X Scrape X Scrape; (facultative N. kennedyi Coll-gather) Rithrogena X Coll-gather * Leptophlebiidae Paraleptophlebia X Coll-gather X Coll-gather; P. zayante (facultative Shred- detritivore) Siphlonuridae * X Coll-gather; Siphlonurus occidentalis (facultative scrape, pred) Leptohyphidae X Coll-gather (=Tricorythidae) Tricorythodes minutus Plecoptera (stoneflies) Chloroperlidae X X Pred; Kathoperla X Coll-gather X Coll-gather; (facultative scrape) Bisancora X Pred? Plumiperla X Pred? Sweltsa X Pred; (facultative Coll-gather) Capniidae X Shred X Shred-detrit Mesocapnia Leuctridae X Shred X Shred-detrit Despaxia augusta X Shred-detrit Paraleuctra sara X Shred-detrit Nemouridae X Shred X Shred-detrit; (facultative Coll- gather) Malenka X Shred X Shred-detrit? M. californica X Nemoura spiniloba X Shred-detrit; (facultative Shred- herb?)

10 Soyedina X Shred X Shred-detrit? Zapada X Shred * Peltoperlidae X Shred X Shred-detrit Soliperla quadrispinula X Perlidae X Pred X Pred Calineuria californica X Pred X Pred Hesperoperla X Pred X Pred H. pacifica Claasenia sabulosa X Pred Perlodidae X Pred X Pred; (facultative Scrape, Pred) Isoperla X Pred; (facultative Coll-gather) Pteronarcyidae X Shred * Pteronarcys Odonata (dragon- and damselflies) Corduligasteridae Corduligaster dorsalis X Pred X Pred Gomphidae Octogomphus specularis X Pred X Pred Aeshnidae * X Pred Megaloptera (dobsonflies, alderflies) Sialidae Sialis californica X Pred X Pred Corydalidae Dysmicohermes X Pred * Trichoptera (caddisflies) Apataniidae Apatania X Scrape X Scrape; (facultative Coll-gather) Brachycentridae Micrasema X Shred * Calamoceratidae Heterplectron X Shred X Shred-detrit califonicum (including wood) Glossosmatidae Glossosoma X Scrape X Scrape Hydropsychidae Hydropsyche X Coll-filt X Coll-filt Parapsyche X Coll-filt * Hydroptilidae Hydroptila X Piercers X Piercers (macrophytes) (macrophytes)

11 Palaeagapetus X Shred * Goeridae Goeracea X Scrape Lepidostomatidae Lepidostoma X Shred X Shred-detrit Limnephilidae Cryptochia X Scrape X Shred-detrit; (facultative Scrape) Ecclisomyia X Coll-gather X Coll-gather; (facultative Scrape- mineral case, Shred-detrit- organic case) Hydatophylax hesperus X Shred X Shred-detrit Psychoglypha X Shred X Shred-detrit subborialis Dicomoecus atripes * X Scrape Onocosmoecus unicolor * X Shred-detrit Odontoceridae Parthina vierra X Shred X Shred-detrit Namamyia plutonis X Coll-gather * Nerophilus californicus X Shred * Philopotamidae Wormaldia X Coll-gather X Phryganeidae Yphnia californica X Pred * Polycentropodidae Polycentropus X Pred X Coll-filt; (facultative Pred, Shred-herb) Rhyocophilidae Rhyacophila X Pred X Pred Uenoidae Neophylax splendens X Scrape X Scrape Farula X Scrape * Lepidoptera (Moths) Crambidae * X Shred-herb; (facultative Scrape) Hemiptera (True bugs) Geridae * X Pred Coleoptera (Beetles) Dytiscidae Agabus pandurus X Pred X Pred Rhantus consimilis * X Pred Ilybius * X Pred

12 Oreodytes * X Pred Elmidae Lara avara * X Shred-detrit (including wood) Narpus concolor X Coll-gather X Coll-gather; (larvae) Scrape (adults) Optioservus X Scrape * quadrimaculatu O. seriatus * X Coll-gather; (larvae) Scrape (adults) Zaitzevia parvula X Coll-gather * Hydraenidae Hydraena X Scrape * Hydrophilidae X Pred (larvae) Ametor scabrosus * X Pred (larvae); Coll- gather (adults) Cymbiodyta * X Pred (larvae); Coll- gather (adults) Hydrochus variolatus * X Shred-herb Psephenidae Acneus X Scrape X Scrape Eubrianax edwardsii X Scrape X Scrape Haliplidae Brychius pacificus * X Shred-herb; (facultative Pred) (larvae and adults) Diptera (true flies) Ceratopogonidae X Pred Atrichopogon * X Coll-gather; (facultative Scrape) Forcipomyia * X Coll-gather; (facultative Scrape) Probezzia * X Pred Leptoconops?? * X Pred Chironomidae X Coll-gather Tanypodinae Paramerina * X Pred? Chironominae Rheotanytarsus * X Coll-filt Tanytarsus * X Coll-filt; (facultative Coll- gather, Scrape) Constempellina * X Coll-gather;

13 (facultative Coll- filt) Dixidae Dixa X Coll-gather X Coll-filt; (facultative Coll- gather) Meringodixa X Coll-gather * Culicidae X Coll-filt * Empididae Chelifera X Pred X Pred Clinocera X Pred * Hemerodromia X Pred * Dolichopodidae * X Pred Ephydridae X Coll-gather Pelina * X Scrape Pelecorhynchidae Glutops X Pred X Pred; (facultative Shred-herb) Psychodie Maruina X Scrape X Scrape; (Coll- gather) Pericoma X Coll-gather * Psychoda * X Coll-gather Ptychopteridae Ptychoptera X Coll-gather X Coll-gather Simuliidae Simulium X Coll-gather X Statiomyidae X Coll-gather * Tabanidae X Pred * Tipulidae Dicranota X Pred X Pred Hexatoma X Pred X Pred Tipula X Shred X Shred-detrit Antocha X Coll-gather * Limnophila X Pred * Limonia X Shred * Pedicia X Pred * Rhabdomastix X Shred * Phoridae * X Coll-gather; (facultative Pred) Tantderidae Prototanyderus * X Pred?

Non-Insects Annelida, Oligochaeta X Coll-gather X Coll-gather

14 Coelenterata, Hydra X Pred X Pred Nematomorpha X Parasite * Hydracarina X Pred X Pred Nematoda X Pred * Turbellaria Planaeidae X Pred X Pred Crustacea Cyclopoida X Coll-gather * Coll-filt Harpactacioda * X Coll-gather Cladocera X Coll-filt * Isopoda Asellus * X Shred-detrit Ostracoda X Coll-gather * Gastropoda Pisidium * X Coll-filt Total taxa 100 102 Missing taxa 48 31

Collector-gatherers 27 27 Predator 27 34 Shredder 18 -- Shredder-detritivore -- 20 Shredder-herbivore -- 3 Scraper 17 15 Collector-filterers 4 7 Piercer (macrophytes) 1 1 Parasite 1 0

15 Appendix 2. Caspar Creek Taxon List Aquatic Insects: Coleoptera Dytiscidae Agabus pandurus Rhantus consimilis Oreodytes Ilybius Elmidae Lara avara Narpus concolor Optioservus seriatus Gyrinidae Gyrinus Haliplidae Brychius pacificus Hydrochidae Hydrochus variolatus Hydrophilidae Ametor scabrosus Cymbiodyta Psephenidae Eubrianax edwardsii Acneus Diptera Ceratopogonidae Atrichopogon Forcipomyia Leptoconops ? Probezzia Chironomidae Tanypodinae Pentaneurini Paramerina Chironominae Tanytarsini Rheotanytarsus Tanytarsus Constempellina Dixidae Dixa Dolichopodidae Empididae

16 Chelifera? Ephydridae Pelina Pelicorhynchidae Glutops Phoridae Diplonevra Protanyderidae ? Protanyderus ? Psychodidae Maruina Psychoda Ptychopteridae Ptychoptera Simuliidae Simulium Tipulidae Dicranota Hexatoma Tipula Ephemeroptera Ameletidae Ameletus sp. Ameletus cooki Baetidae Baetis bicaudatus Baetis tricaudatus Diphetor hageni Fallceon quilleri Ephemerellidae Drunella flavilinea Ephemerella dorothea infrequens Serratella teresa Serratella tibialis Timpanoga hecuba pacifica Heptageniidae Cinygma Cinygmula Epeorus Ironodes Nixe kennedyi Leptophlebiidae Paraleptophlebia spp. Paralpetophlebia zayante

17 Siphlonuridae Siphlonurus occidentalis Leptohyphidae (=Tricorythidae) Tricorythodes minutus Hemiptera Gerridae Metrobates trux Veliidae Microvelia californiensis Lepidoptera Geometridae Pyralidae Megaloptera Sialidae Sialis Odonata Aeshnidae Cordulegastridae Cordulegaster dorsalis Gomphidae Octogomphus specularis Plecoptera Chloroperlidae Bisancora pastina? Kathroperla Plumiperla Sweltsa Capniidae Mesocapnia Leuctridae Despaxia augusta Paraleuctra sara Nemouridae Malenka californica Nemoura spiniloba Soyedina Peltoperlidae Perlidae Calineuria californica Claasenia sabulosa Hesperoperla pacifica Perlodidae Isoperla Trichoptera

18 Apataniidae Apatania Calamoceratidae Heteroplectron californicum Glossosomatidae Glossosoma Hydropsychidae Hydropsyche Hydroptilidae Hydroptila Lepidostomatidae Lepidostoma Limnephilidae Hydatophylax hesperus Psychoglypha sp. Psychoglypha subborealis Dicosmoecus atripes Onocosmoecus unicolor Ecclisomyia Cryptochia Odontoceridae Parthina vierra Philopotamidae Wormaldia Polycentropodidae Polycentropus Rhyacophilidae Rhyacophila Uenoidae Neophylax splendens Other Aquatic Invertebrates: Acari (subclass) Anthoathecatae

Hydridae Hydra Basommatomorpha Planorbidae Collembola Entomobryidae Hypogastruridae Simithuridae Dicyrtoma Tomoceridae

19 Tomocerus Cyclopoida Harpacticoida Isopoda Asellidae Asellus Nemata (phylum) Oligochaeta (subclass) Poecilostomatoida Ergasilidae Ergasilus Tardigrada (phylum) Turbellaria (class) Veneroida Pisidiidae Pisidium Terrestrial Insects: Coleoptera Corylophidae Curculionidae Staphylinidae

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Diptera Cecidomyiidae Sciaridae Xylophagidae? Hymenoptera Apidae Formicidae Mymaridae Pteromalidae Scelionidae Thysanoptera Thripidae Other Taxa: Araneae Pseudoscorpiones

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